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Article history:
Received 12 October 2013
Received in revised form 22 January 2015
Accepted 23 January 2015
Keywords:
Priority index
Ecological oddity
Phylogenetic distinctness
Extrinsic factors
Pseudoboini
Neotropical snakes
a b s t r a c t
Species differ in their need for conservation action and in their relative importance for conserving current
and historic ecological and evolutionary diversity. Given the present biodiversity crisis and the lack of
resources, threatened species must be differentiated from each other so that those presenting higher
conservation priority can be attended rst. Here we propose a novel approach to calculate a priority
index (PI) for species within monophyletic groups, by combining life history traits, extrinsic factors,
ecological singularity, and phylogenetic distinctness. To test our approach we used a group of Neotropical
snakes, the pseudoboines, as our model lineage. To create the PI we combined four different indices:
intrinsic vulnerability to extinction (IVE, comprised by six factors), extrinsic vulnerability to extinction
(EVE, comprised by three factors), ecological oddity (EO, four factors) and phylogenetic distinctness (PD).
Intrinsic vulnerability to extinction was evenly distributed across the clade and EVE was higher in species
present in the Brazilian hotspots of Biodiversity, Atlantic Forest and Cerrado. As expected due to the
nature of the index, a few species that differ from the average phenotype presented high EO values,
whereas PD values did not vary greatly among pseudoboines. Representatives from almost all clades
within the pseudoboines appear among the ten highest PI values, maximizing the phylogenetic diversity
of the prioritized taxa. Although it is not possible to compare values obtained in studies to different
lineages (indices are clade-specic), extending this approach to more inclusive lineages (e.g., families)
might enhance the quality of future prioritization processes. The method we propose would be especially
useful for taxonomically driven conservation action plans.
2015 Elsevier GmbH. All rights reserved.
Introduction
Given the biodiversity crisis and the relative scarcity of
resources (Barnosky et al. 2011; Brooks et al. 2006) threatened
species must be differentiated from each other so that those
presenting higher conservation priority can be attended to rst
(Wilson et al. 2009). Thus, a major challenge for conservation
focused on species is to set priorities for conservation efforts (Pimm
et al. 2001; Wilson et al. 2006). Although the prioritization of conservation efforts may involve several factors such as socioeconomic
and political issues (Eklund et al. 2011; Polasky 2008; Wilson et al.
2009; Wilson et al. 2011), having the knowledge of the vulnerability to extinction of a species can help to predict the outcomes
Corresponding author. Tel.: +55 11 3091 7600; fax: +55 11 3091 8096.
E-mail address: gaiarsa.mp@gmail.com (M.P. Gaiarsa).
http://dx.doi.org/10.1016/j.jnc.2015.01.006
1617-1381/ 2015 Elsevier GmbH. All rights reserved.
50
on lizards and small mammals, whereas some species are specialized in other prey types (e.g., Rhachidelus brazili feeds on bird
eggs; Gaiarsa et al. 2013). Additionally, the tribe is composed
predominantly of terrestrial species (e.g., Clelia spp. and Boiruna
spp.), although some species are semi-arboreal (e.g., Drepanoides
anomalus and Siphlophis cervinus) or semi-fossorial (Phimophis spp.;
Gaiarsa et al. 2013). Hence, this tribe is an ideal model for this
study because of its diversity of ecological features and the relative wealth of natural history data (Gaiarsa et al. 2013). Full details
of the data set and sources are available elsewhere (Tables A1 and
A2; Alencar et al. 2013; Gaiarsa et al. 2013). We followed the taxonomy of Grazziotin et al. (2012) and Zaher et al. (2009) and made
no distinction among subspecies.
Vulnerability to extinction
We adapted the method of Millsap et al. (1990) using data on
factors known to affect species survival to create two indices of
vulnerability to extinction: intrinsic and extrinsic vulnerability to
extinction (IVE and EVE, respectively; see below). We only included
in the analysis species for which data was available for at least 50%
of the factors used to create each index.
Intrinsic vulnerability to extinction (IVE)
We selected six factors that are related to characteristics that
could decrease a species ability to cope with negative alterations
in its habitat, and that are either related to narrow niches and/or to
low abundance (Mckinney 1997): (i) body size, (ii) mean fecundity,
(iii) dietary specialization, (iv) geographic distribution, (v) elevational range and (vi) ability to persist in altered habitats. We initially
included habitat breadth (HB, measured as the number of Terrestrial Ecoregions of the World in which the species occurs, Olson et al.
2001, encompassed by a species geographic range) as a seventh
factor. However, preliminary analyses indicated a high correlation
between HB and the factor geographic distribution (GD; r = 0.81,
Table A3; see below for information on how GD was quantied), so
we excluded habitat breadth from IVE calculations. The remaining
factors had a relatively low correlation (r < 0.45 in all cases, Table
A3). Factors were as follows:
1. Body size (BS): larger species tend to be more vulnerable to
extinction because they are in general less abundant, have later
sexual maturity and are more long-living, thus being less able
to recover from population declines (e.g., Foufopoulos & Ives
1999; Mckinney 1997; Purvis et al. 2000). We used the maximum known snout-vent length for each species, regardless of
the sex.
2. Mean fecundity (MF): low fecundity populations tend to be more
prone to extinction because they take longer to recover from
declines than do high fecundity populations (e.g., Mckinney
1997; Pimm et al. 1988; Purvis et al. 2000). We used mean clutch
size, regardless of the number of litters available.
3. Dietary specialization (DS): animals that are specialists in
resource use (e.g., prey, habitat) tend to be less able to cope with
changes in the resource base (either by anthropogenic or natural causes) and thus, are more vulnerable to extinction (e.g.,
Mckinney 1997; Purvis et al. 2000). To characterize the degree
of diet specialization we used the percentage of the most important prey item in the diet. Thus, the greater the contribution of
one type of prey to the diet, the more specialized in one prey
the species is. Prey items considered were: amphibians, lizards,
lizard eggs, snakes, birds, bird eggs, and mammals. We considered every record, regardless of the number of individual prey
available for each species.
4. Geographic distribution (GD): the smaller the geographic distribution the more vulnerable to extinction a species tends to be
(see, e.g., Cardillo et al. 2008; Fisher & Owens 2004; Purvis et al.
2000). We used occurrence points from museum records and
literature data to estimate the geographic distribution of each
species (for a discussion on different methods see Attorre et al.
2013). Because good distribution maps are not available for most
snakes (see, e.g., Uetz 2013), and very few were available for our
model lineage, GD was calculated according to the available data
(for details on how GD was calculated for pseudoboine snakes,
see supplementary material). Our geographic distribution factor is equivalent to the IUCNs extent of occurrence (EOO; IUCN
2001).
5. Elevational range (EL): animals occurring in a restricted vertical distribution may be more vulnerable to extinction because
they tend to be stenothermics and stenobarics, and thus less
able to cope with habitat change (either by anthropogenic or
natural causes; e.g., Mckinney 1997). We assessed this factor by
calculating the elevational range within the species geographic
distribution (with the same database used to calculate the factor
Geographic Distribution).
6. Ability to persist in altered habitats (AAH; adapted from Filippi
& Luiselli 2000): vulnerability to extinction tends to be greater
in animals that are less able to persist in altered habitats (e.g.,
Fisher & Owens 2004; Purvis et al. 2000). This factor was assessed
based on the personal experience of researchers. Experienced
researchers were asked to assign values for each species as follows: 05 for species which are able to persist in disturbed
habitats (including urban areas); 611 for those which may occasionally persist in disturbed habitats (found in rural areas where
small patches of natural vegetation are available); 1217 for
species which rarely persist in disturbed habitats (may be found
in habitat patches); and 1823 for species which do not persist
in disturbed habitats (found only in large extensions of natural habitat). We employed the mean of the scores given by each
expert for each species. Even though in some cases the opinion
of researchers varied considerably (probably due to the subjectivity involved, see e.g., Keith et al. 2004; Regan et al. 2004, but
also because of geographical variation in this feature), we believe
this metric is still an informative measure of species sensibility
to human disturbance.
51
52
of the variable dietary breadth. We only considered prey items representing over 20% of the diet. The mean of these four factors was
then ranked, such that the higher the score, the greater the species
rank and when ties occurred we employed the mean ranking
score.
Phylogenetic distinctness
We used the most recent published phylogenetic hypothesis
for the tribe Pseudoboini (Grazziotin et al. 2012; Fig. A1). We also
included species that were not in the original phylogeny considering their afnities with the species that were already included
(see, e.g., Martins et al. 2001) using literature information (e.g.,
Vidal et al. 2010; Zaher et al. 2009; Fig. A1) and expert knowledge (H. Zaher, pers. comm.). We considered Mays distinctness
measure (May 1990) as our phylogenetic distinctness (PD) scores,
which measures the sum of the number of descendants of nodes
Priority index
Although the indices (IVE, EVE, EO, and PD) used to calculate
the priority index (PI) have different magnitude, when we used the
rankings of each species they became standardized, rendering them
comparable. We then tested for correlation among indices using a
Pearson rank correlation test (after testing indices for normality;
Table A5). We only included in the analysis species for which at
least three of the indices were available.
Table 1
Ranking of factors scores used to build the index of intrinsic (IVE) and extrinsic (EVE) vulnerability to extinction and the values for ecological oddity (EO) and Phylogenetic
Distinctness (PD) for the species of the tribe Pseudoboini (refer to text for further details). Species appear in alphabetical order. BS: body size; MF: mean fecundity; DS:
dietary specialization; GD: geographic distribution; EL: elevational range; AAH: ability to persist in altered habitats; RH: percentage of remaining habitats; PA: percentage of
protected areas along the distribution; HI: mean human inuence index along the distribution; HB: habitat breadth; DB: dietary breadth. Variables for which data was not
available are indicated with . Habitat breadth and DF were excluded from the calculation of IVE and EVE, respectively (refer to text for further details).
Species
Boiruna maculata
Boiruna sertaneja
Clelia clelia
Clelia equatoriana
Clelia errabunda
Clelia hussami
Clelia langeri
Clelia plumbea
Clelia scytalina
Drepanoides anomalus
Mussurana bicolor
Mussurana montana
Mussurana quimi
Oxyrhopus clathratus
Oxyrhopus doliatus
Oxyrhopus erdisii
Oxyrhopus tzingeri
Oxyrhopus formosus
Oxyrhopus guibei
Oxyrhopus leucomelas
Oxyrhopus marcapatae
Oxyrhopus melanogenys
Oxyrhopus occipitalis
Oxyrhopus petola
Oxyrhopus rhombifer
Oxyrhopus trigeminus
Oxyrhopus vanidicus
Paraphimophis rusticus
Phimophis guerini
Phimophis guianensis
Phimophis vittatus
Pseudoboa coronata
Pseudoboa haasi
Pseudoboa martinsi
Pseudoboa neuwiedii
Pseudoboa nigra
Pseudoboa serrana
Rhachidelus brazili
Rodriguesophis chui
Rodriguesophis iglesiasi
Rodriguesophis scriptorcibatus
Siphlophis cervinus
Siphlophis compressus
Siphlophis leucocephalus
Siphlophis longicaudatus
Siphlophis pulcher
Siphlophis worontzowi
IVE
EVE
EO
PD
BS
MF
DS
GD
EL
AAH
RH
PA
HI
BS
MF
HB
DB
39
40
41
37
36
23.5
34
42
29
7
11
17
22
28
3
23.5
14
13
27
18
12
15
38
21
5
6
26
33
25
19
32
31
35
2
4
1
20
30
8
16
10
9
13
8
2
29
9.5
9.5
5
15
6
27.5
16
12
14
3
11
24
22
25
21
18.5
20
7
26
30
23
17
18.5
27.5
12
15
4.5
30.5
18
20
8
8
16.5
23
19
3
30.5
1
4.5
11
8
8
26
2
13
30.5
8
14
21
30.5
30.5
22
27
30.5
16.5
24.5
24.5
13
17
2
34
41
39
4
30
8
27
35
15
31
38
16
29
36
11
21
10
3
6
18
20
9
24
23
1
32
14
22
12
40
28
19
7
5
33
25
37
26
21
31.5
10
7
41
8
23
9
38
39
17.5
15
29
28
20
2
3
40
12
5
14
30
11
6
17.5
37
4
35
33
22
24
25
17.5
26
34
1
36
31.5
17.5
27
13
10
30
27
21
23
13
30
24.5
1
3
17
7.5
19
14
7.5
12
24.5
2
5
16
11
27
22
30
18
20
6
15
27
27
37
38
26
40
19
24
10
23
14
25
5
29
1
28
21
7
13
6
34
33
12
11
39
36
30
9
4
8
17
2
22
16
35
15
18
31
32
20
3
35
34
18
15
40
2
20
38
9
39
23
30
22
41
29
16
1
12
14
21
32
24
5
36
26
17
37
7
31
4
11
28
3
33
19
10
13
25
27
6
8
26
23
11
31
27
14
20
37
3
15
41
34
32
25
36
24
10
6
9
19
22
18
2
30
21
13
17
4
33
1
12
29
38
39
16
5
7
28
35
40
8
0.290
0.294
0.386
0.153
0.146
0.040
0.119
0.452
0.082
0.173
0.077
0.020
0.029
0.060
0.431
0.040
0.039
0.041
0.050
0.012
0.059
0.036
0.274
0.023
0.181
0.180
0.045
0.118
0.044
0.006
0.107
0.101
0.144
0.516
0.346
0.602
0.002
0.096
0.143
0.024
0.089
0.121
0.068
0.135
0.269
0.269
0.463
0.123
0.123
0.220
0.060
0.260
0.229
0.345
0.022
0.086
0.068
0.248
0.117
0.155
0.132
0.170
0.102
0.053
0.081
0.146
0.178
0.530
0.143
0.039
0.053
0.229
0.271
0.127
0.734
0.185
1.030
0.252
0.201
0.116
0.201
0.075
0.252
0.185
0.127
0.729
0.174
0.127
0.127
0.116
0.146
0.678
0.249
0.293
0.049
0.174
0.049
0.116
0.075
0.332
0.252
0.226
0.226
0.313
0.553
0.428
0.185
0.417
0.332
0.127
0.127
0.428
0.030
0.231
0.092
0.066
0.077
0.085
0.787
0.500
0.077
0.058
0.063
0.068
0.056
0.043
0.288
0.059
0.050
0.060
0.083
0.040
0.212
0.046
0.111
0.069
0.029
0.800
0.043
0.043
0.035
0.042
0.043
0.060
0.036
0.036
1.333
1.333
1.143
1.143
1.143
1.143
1.143
1.600
1.143
1.143
1.143
1.600
1.143
1.333
1.143
1.143
1.333
1.333
1.333
1.143
1.600
1.231
1.231
1.231
1.143
1.000
1.067
1.067
1.000
1.067
1.600
1.778
1.778
1.778
2.286
2.286
2.286
2.286
2.286
2.286
53
Results
Discussion
There was great variation in the biological features for the group
studied (Table A1; Gaiarsa et al. 2013). In general, all the species
with the highest IVE presented high values for the factors GD and
BS (Table 1). The index of extrinsic vulnerability to extinction (EVE)
was calculated for 41 species and ranged from 4.3 to 35.7, with a
mean of 20.7 8.8 (Table 2). The index of intrinsic vulnerability to
extinction (IVE) was calculated for 39 species and ranged from 10.3
to 35.2, with a mean of 19.4 5.3 (Table 2).
Ecological oddity (EO) was calculated for 39 species and ranged
from 0.06 to 0.53 (Table 2), with a mean of 0.18 0.11, and
phylogenetic distinctness (PD) was obtained for 40 species. Due
to polytomies in our phylogeny, all the species from the genus
Siphlophis presented the highest PD (2.29), followed by the genus
Rodriguesophis (Table 1 and Fig. A1). Finally, priority index (PI)
was calculated for 39 species and ranged from 5.38 to 30.46
(Table 2).
Table 2
Mean ranking scores for all the indices: intrinsic vulnerability to extinction (IVE), extrinsic vulnerability to extinction (EVE), ecological oddity (EO), phylogenetic distinctiveness
(PD) and priority index (PI) for the species of the tribe Pseudoboini. See text for further detail. Species are ranked in descending order according to their priority index. We
only included in the analysis species for which at least three of the indices were available (Clelia errabunda, Oxyrhopus doliatus, O. erdisii, O. tzingeri, O. formosus, O. leucomelas,
O. marcapatae and Rodriguesophis chui were not evaluated). Missing values are indicated with .
Species
IVE
EVE
PD
EO
PI
Clelia hussami
Rhachidelus brazili
Rodriguesophis scriptorcibatus
Rodriguesophis iglesiasi
Siphlophis pulcher
Boiruna sertaneja
Boiruna maculata
Siphlophis leucocephalus
Paraphimophis rusticus
Drepanoides anomalus
Clelia equatoriana
Siphlophis compressus
Siphlophis longicaudatus
Pseudoboa serrana
Siphlophis cervinus
Clelia clelia
Phimophis guianensis
Mussurana bicolor
Oxyrhopus petola
Clelia plumbea
Oxyrhopus guibei
Phimophis vittatus
Clelia langeri
Oxyrhopus clathratus
Oxyrhopus trigeminus
Clelia scytalina
Oxyrhopus rhombifer
Mussurana montana
Mussurana quimi
Phimophis guerini
Pseudoboa haasi
Siphlophis worontzowi
Oxyrhopus occipitalis
Pseudoboa coronata
Pseudoboa nigra
Oxyrhopus melanogenys
Pseudoboa martinsi
Pseudoboa neuwiedii
Oxyrhopus vanidicus
Clelia errabunda
Oxyrhopus doliatus
Oxyrhopus erdisii
Oxyrhopus tzingeri
Oxyrhopus formosus
Oxyrhopus leucomelas
Oxyrhopus marcapatae
Rodriguesophis chui
35.17
24.5
20.5
23.25
23.5
23.58
18
25.75
17
20.83
26
22.5
16.58
26.13
15.17
14.42
22
17.92
10.33
18.17
13.92
13.75
27.88
22.5
13.42
22.67
11.42
19.5
14.7
18.58
24
19.9
20.8
16.08
15.83
13
22.42
15.83
11
35.67
29.33
23.33
22
31.33
29.33
28
25.67
7.33
24
12.67
31.33
21
10
22.33
22
25.67
15.33
19.67
22
28
8
19.67
25
33
29.33
26
29.67
28.67
22.67
6.33
12
6.67
19.67
8.33
4.33
13.33
6.33
31.67
22.67
10.67
12
29.5
33
33
37.5
24.5
24.5
37.5
29.5
29.5
12
37.5
37.5
4
37.5
12
20
12
24.5
12
24.5
20
29.5
24.5
12
24.5
12
12
20
1.5
37.5
12
12
1.5
12
4
4
12
12
33
39
37
34
33
29
24
30
9
25
38
26
15
2
35
23
36
20
28
32
31
18
16
22
4
12
7
8
13
14
3
17
1
19
27
21
11
10
5
6
30.46
30.08
29.17
28.15
28.00
25.85
25.46
25.06
24.29
23.92
22.00
21.92
21.85
21.53
21.42
21.19
21.00
20.90
20.54
20.21
19.61
19.44
19.29
18.92
18.73
18.67
18.31
17.63
17.59
17.56
16.29
16.18
15.95
15.44
14.50
11.08
10.19
9.54
8.83
54
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